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Cited by 287Spatial Cognition and NavigationMemory and Neural MechanismsVestibular and auditory disorders

Anchoring the neural compass: coding of local spatial reference frames in human medial parietal lobe

Open access
Marchette, Steven A, Vass, Lindsay K, Ryan, Jack, Epstein, Russell A·2014·Nature Neuroscience
DOI 10.1038/nn.3834PMC PMC4309016
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Abstract

The neural systems that code for location and facing direction during spatial navigation have been investigated extensively; however, the mechanisms by which these quantities are referenced to external features of the world are not well understood. To address this issue, we examined behavioral priming and functional magnetic resonance imaging activity patterns while human subjects recalled spatial views from a recently learned virtual environment. Behavioral results indicated that imagined location and facing direction were represented during this task, and multivoxel pattern analyses indicated that the retrosplenial complex (RSC) was the anatomical locus of these spatial codes. Critically, in both cases, location and direction were defined on the basis of fixed elements of the local environment and generalized across geometrically similar local environments. These results suggest that RSC anchors internal spatial representations to local topographical features, thus allowing us to stay oriented while we navigate and retrieve from memory the experience of being in a particular place.

AI snapshot

Written from the paper’s full text.

TL;DR

Using behavioral priming and fMRI multivoxel pattern analysis during a judgment of relative direction task, the authors show that human retrosplenial complex (RSC) codes imagined facing direction and location. These spatial codes are anchored to local environmental geometry and generalize across geometrically similar local environments rather than using a global reference frame.

Key findings

  • Behavioral priming in Experiment 1 showed faster reaction times when imagined direction repeated (3.31 ± 0.20 s same vs. 3.56 ± 0.18 s different; F(1,21) = 11.631, p = 0.003) and when imagined location repeated (3.31 ± 0.20 s same vs. 3.56 ± 0.18 s different; F(1,21) = 5.695, p = 0.026), with no interaction between direction/location and museum.
  • In RSC, multivoxel patterns were more similar for views facing the same local direction (F(1,23) = 20.009, p = 0.00017) and for views in the same or geometrically equivalent local location (F(1,23) = 7.162, p = 0.013), with no global direction or global location coding.
  • RSC pattern similarities reconstructed within-museum spatial relationships significantly better than chance (D: 16%, p = 0.00008), and across-museum reconstruction was also significant (D: 47%, p = 0.015); other ROIs did not reliably reconstruct space (all Ds > 58%, ps > 0.1).
  • Whole-brain searchlight found local direction decoding in bilateral RSC and left superior parietal lobule; local location coding was not significant at corrected levels, and no global direction coding was observed even at liberal thresholds.

Methods

  • Two experiments used a virtual park containing four visually distinct “museums” with identical internal geometry but different global orientations; participants learned object locations and performed a judgment of relative direction task requiring imagined location and facing direction.
  • Experiment 1 tested 22 subjects behaviorally; Experiment 2 scanned 24 subjects with fMRI on a 3T Siemens Trio while they performed a modified JRD task.
  • Analyses included repeated-measures ANOVAs and paired t-tests on reaction times and multivoxel pattern similarities, Pearson correlations between activity patterns across independent carryover sequences, multidimensional scaling/Procrustes reconstruction, and whole-brain searchlight with Monte Carlo correction.

Limitations

  • The paper states that “local” is caveated because reference and target objects were always in the same museum; if targets outside the museum had been used, subjects might have accessed a global frame.
  • The paper notes that all museums were learned simultaneously, which may have encouraged a single spatial frame or schema, and reports anecdotal evidence that subjects sometimes traveled to the right location in the wrong museum.
  • The paper states that virtual navigation lacked proprioceptive and vestibular cues, that a different training regime might reduce cross-environment generalizability, and that it remains unclear whether walls, the single door, or the principal axis defined heading.
  • Evident from the text: Experiment 2 did not show significant behavioral direction or location priming (direction: F(1,23) = 1.501, p = 0.233; location: F(1,23) = 1.190, p = 0.287), which the authors attribute to a longer intertrial interval; normality of data distributions was assumed but not formally tested.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1038/nn.3834

Cites

61 of 61 references verified0 citation errors · 0 not checkable

Statistics

32 tests recomputed · 0 with the reported and recomputed p on opposite sides of .05

Claims the paper makes

0 of 55 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    In Experiment 1, there was no significant interaction between direction and museum.

    “No significant interaction between direction and museum was observed, F(1,21) = 0.734, p = 0.401.”
    Quote in the paperSubstantial claim
  • Findingown data

    In Experiment 1, a repeated-measures ANOVA revealed a main effect of direction, with faster reaction times when imagined direction was repeated over successive trials (3.31 ± 0.20 s same vs. 3.56 ± 0.18 s different).

    “A repeated-measures ANOVA with direction (same/different as preceding trial) and museum (same/different as preceding trial) as factors revealed a main effect of direction, F(1,21) = 11.631, p = 0.003, with faster reaction times when imagined direction was repeated over successive trials (3.31 ± 0.20 s same vs. 3.56 ± 0.18 s different) (Fig. 3a, left panel).”
    Quote in the paperSubstantial claim
  • Findingown data

    In Experiment 1, there was a main effect of repeating museum, with faster reaction times when reference objects on successive trials were located within the same museum (3.27 ± 0.19 s same vs. 3.59 ± 0.19 s different).

    “We also observed a main effect of repeating museum, F(1,21) = 11.802, p = 0.002, with faster reaction times when reference objects on successive trials were located within the same museum (3.27 ± 0.19 s same vs. 3.59 ± 0.19 s different).”
    Quote in the paperSubstantial claim
  • Findingown data

    Participants in Experiment 1 responded accurately on most trials (M = 89.8% correct, S = 6%).

    “Participants responded accurately on most trials (M = 89.8% correct, S = 6%).”
    Quote in the paperSubstantial claim
  • Findingown data

    RSC coded imagined facing direction.

    “The results indicated that RSC coded imagined facing direction (Fig. 4a, left panel).”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. 34
  • May combine two claims. 38

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    There was a marginal effect of repeating museum in the location analysis.

    “There was a marginal effect of repeating museum, F(1,21) = 4.227, p = 0.052.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    Directional and location priming effects were not significant in Experiment 2.

    “Directional and location priming effects were not significant in Experiment 2 (direction: F(1,23) = 1.501, p = 0.233, n.s.; location: F(1,23) = 1.190, p = 0.287, n.s.)”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    There was no effect of museum identity and no interaction between direction and museum in the RSC direction analysis.

    “There was no effect of museum identity, F(1,23) = 2.450, p = 0.131 and no interaction between direction and museum, F(1,23) = 0.005, p = 0.942.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    In the controlled direction analysis, there was no main effect of museum and no interaction of direction and museum.

    “with no main effect of museum, F(1,23) = 0.850, p = 0.366, and no interaction of direction and museum, F(1,23) = 1.043, p = 0.318.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    The starting view was maximally similar to itself (scaled to 1.0), indicating consistency of the response pattern across the first and second half of the experiment.

    “The starting view was maximally similar to itself (scaled to 1.0), indicating consistency of the response pattern across the first and second half of the experiment.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    There was no main effect of museum and no interaction of location with museum in the RSC location analysis.

    “There was no main effect of museum, F(1,23) = 2.719, p = 0.113 and no interaction of location with museum F(1,23) = 0.284, p = 0.600.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    In the 90-degree controlled location analysis, there was no main effect of museum and no interaction of location by museum.

    “and no main effect of museum, F(1,23) = 1.834, p = 0.189 and no interaction of location by museum F(1,23) = 0.768, p = 0.390.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    Across museums, the most similar views were the locally analogous view and the view facing the same local direction; there was also some similarity for the view facing the orthogonal direction in the equivalent location.

    “Across museums, the most similar views were the locally analogous view and the view facing the same local direction; there was also some similarity for the view facing the orthogonal direction in the equivalent location.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    An analogous searchlight analysis for local location coding found no brain regions at corrected significance levels, although coding in bilateral RSC, left SPL, and other regions of posterior parietal cortex were observed at uncorrected significance levels.

    “An analogous searchlight analysis for local location coding found no brain regions at corrected significance levels, although coding in bilateral RSC, left SPL, and other regions of posterior parietal cortex were observed at uncorrected significance levels.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    We also observed location-dependent priming and location-dependent activity patterns.

    “Moreover, we also observed location-dependent priming and location-dependent”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    We observed orientation-dependent priming and orientation-dependent activity patterns when orientation was defined relative to local geometry.

    “Consistent with these previous results, we observed orientation-dependent priming and orientation-dependent activity patterns when orientation was defined relative to local geometry.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    During pre-scan encoding, subjects sometimes traveled to the right location in the wrong museum.

    “when subjects were asked to find an object, they sometimes traveled to the right location in the wrong museum.”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    In rodents, location information is encoded by place and grid cells in the hippocampal formation, and directional information is coded by head direction cells in Papez circuit structures.

    “In rodents, location information is encoded by place and grid cells in the hippocampal formation,1, 2 and directional information is coded by head direction (HD) cells in Papez circuit structures3”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [1] O'Keefe, J. & Dostrovsky, J. The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat. Brain Res. 34, 171–175 (1971). source

      Not checked in the free run.

    • cites [2] Hafting, T., Fyhn, M., Molden, S., Moser, M.-B. & Moser, E.I. Microstructure of a spatial map in the entorhinal cortex. Nature 436, 801–806 (2005). source

      Not checked in the free run.

    • cites [3] Taube, J.S., Muller, R.U. & Ranck, J.B. Head-direction cells recorded from the postsubiculum in freely moving rats. I. Description and quantitative analysis. J. Neurosci. 10, 420–435 (1990). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S4 (based).

    Premisereasoning

    To use local landmarks to anchor sense of direction, one must have a representation of one's heading relative to local environmental features that is at least potentially separable from the internal sense of direction supported by HD cells.

    “To do this, however, one must have a representation of one’s heading relative to local environmental features that is at least potentially separable from the internal sense of direction supported by the HD cells.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Premisecitation

    Previous behavioral work with the JRD task has found evidence for coding of locations and directions in spatial reference frames aligned to local features of the environment such as room geometry or the arrangement of objects within a room, and also in frames aligned to egocentric axes when external features are absent.

    “Previous behavioral work with the JRD task has found evidence for coding of locations and directions in spatial reference frames aligned to local features of the environment such as room geometry or the arrangement of objects within a room (and also in frames aligned to egocentric axes when external features are absent)19, 20.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [19] Shelton, A.L. & McNamara, T.P. Systems of spatial reference in human memory. Cognit. Psychol. 43, 274–310 (2001). source

      Not checked in the free run.

    • cites [20] Mou, W. & McNamara, T.P. Intrinsic frames of reference in spatial memory. J. Exp. Psychol. Learn. Mem. Cogn. 28, 162–170 (2002). source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation

    Recent work has observed similar spatial codes in these regions in humans.

    “recent work has observed similar spatial codes in these regions in humans4–8”
    Quote in the paperSubstantial claim0 of 5 sources checked
    • cites [4] Vass, L.K. & Epstein, R.A. Abstract representations of location and facing direction in the human brain. J. Neurosci. 33, 6133–6142 (2013). source

      Not checked in the free run.

    • cites [5] Hassabis, D. et al. Decoding neuronal ensembles in the human hippocampus. Curr. Biol. 19, 546–554 (2009). source

      Not checked in the free run.

    • cites [6] Doeller, C.F., Barry, C. & Burgess, N. Evidence for grid cells in a human memory network. Nature 463, 657–661 (2010). source

      Not checked in the free run.

    • cites [7] Ekstrom, A.D. et al. Cellular networks underlying human spatial navigation. Nature 425, 184–188 (2003). source

      Not checked in the free run.

    • cites [8] Jacobs, J. et al. Direct recordings of grid-like neuronal activity in human spatial navigation. Nat. Neurosci. 16, 1188–1190 (2013). source

      Not checked in the free run.

  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    A possible solution is to use one's perceived orientation relative to local landmarks to anchor one's sense of direction, which can then be maintained during navigation through idiothetic cues.

    “A possible solution is to use one’s perceived orientation relative to local (currently-visible) landmarks to anchor one’s sense of direction, which can then be maintained during navigation through idiothetic cues10–12.”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [10] Valerio, S. & Taube, J.S. Path integration: how the head direction signal maintains and corrects spatial orientation. Nat. Neurosci. 15, 1445–1453 (2012). source

      Not checked in the free run.

    • cites [11] Yoder, R.M. et al. Both visual and idiothetic cues contribute to head direction cell stability during navigation along complex routes. J. Neurophysiol. 105, 2989–3001 (2011). source

      Not checked in the free run.

    • cites [12] Yoder, R.M., Clark, B.J. & Taube, J.S. Origins of landmark encoding in the brain. Trends Neurosci. 34, 561–571 (2011). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S2 (checkable).

    Premisereasoning

    For a neural compass to be useful, heading must be defined relative to fixed features of the environment.

    “However, for a neural compass to be useful, it is not enough to represent direction in arbitrary coordinates – the heading must be defined relative to fixed features of the environment, just as the heading revealed by a magnetic compass is defined relative to the north-south axis of the earth.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationown data

    The absence of an interaction suggests that direction was defined consistently across all four museums; local North in one museum was equivalent to local North in another, even though these were different directions in the global reference frame.

    “The absence of an interaction is important because it suggests that direction was defined consistently across all four museums. That is, “local North” in one museum was equivalent to “local North” in another, even though these were different directions in the global reference frame.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    HD cells support an internal compass that represents the animal's heading and updates this quantity as the animal moves.

    “This suggests that the HD cells support an internal compass that represents the animal's heading (i.e. facing direction) and updates this quantity as the animal moves.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S4 (based).

    Premisenone given

    In the absence of magnetoception or a sidereal/solar compass, there is no single perceptible feature that consistently indicates direction across all terrestrial environments.

    “This presents a challenge: in the absence of magnetoception or a sidereal/solar compass, there is no single perceptible feature that consistently indicates direction across all terrestrial environments.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationown data

    RSC encodes imagined facing direction during memory retrieval and does so in a way that is referenced to the local environment and is consistent across geometrically-similar environments.

    “These data suggest that RSC encodes imagined facing direction during memory retrieval and does so in a way that is referenced to the local environment and is consistent across geometrically-similar environments.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Location was represented in this task as well as direction and-like direction-it was coded using a locally-anchored reference frame that generalizes across geometrically-equivalent environments.

    “These results showed that location was represented in this task as well as direction and—like direction—it was coded using a locally-anchored reference frame that generalizes across geometrically-equivalent environments.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Participants represented their facing direction during the JRD task and updated this representation when the direction changed from trial to trial, leading to a reaction time cost.

    “These data indicate that participants represented their facing direction during the JRD task and updated this representation when the direction changed from trial to trial, thus leading to an reaction time cost.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).

    Interpretationreasoning

    The faster reaction times in Experiment 2 were likely because fMRI participants received an additional training session on the day of the scan.

    “likely because fMRI participants received an additional training session on the day of the scan”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationown data

    Priming occurs between geometrically-equivalent locations, not only when these locations are physically the same, but also when they are different locations in different museums.

    “indicating that priming occurs between geometrically-equivalent locations, not only when these locations are physically the same, but also when they are different locations in different museums.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Just as direction coding can be observed in RSC when location is completely controlled, location coding can be observed when direction is completely controlled.

    “Thus, just as direction coding can be observed in RSC when location is completely controlled, so can location coding be observed when direction is completely controlled.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Location in RSC is encoded using a local reference frame that is anchored to each museum and generalizes across museums.

    “moreover, like direction, it is encoded using a local reference frame that is anchored to each museum and generalizes across museums.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    RSC coded both direction and location in the local reference frame.

    “Our analyses suggest that RSC coded both direction and location in the local reference frame.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Activity patterns in RSC contain rich information about the implied headings and locations of imagined views that could be used to link views to long-term spatial knowledge, and thus to localize and orient oneself within a mental map of local space.

    “These results provide another demonstration that activity patterns in RSC contain rich information about the implied headings and locations of imagined views that could be used to link views to long-term spatial knowledge, and thus to localize and orient oneself within a mental map of local space.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    This generalization is consistent with the idea that RSC represents views relative to the frame of the local environment and can generalize these relationships to different environments with similar geometry.

    “This generalization is consistent with the idea that RSC represents views relative to the frame of the local environment and can generalize these relationships to different environments with similar geometry.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Both behavioral priming and fMRI signals revealed spatial codes that were anchored to the local environmental features and generalized across geometrically similar local environments, rather than codes that were idiosyncratic to each local environment, or codes that were anchored to a global reference frame.

    “Thus, both behavioral priming and fMRI signals revealed spatial codes that were anchored to the local environmental features and generalized across geometrically similar local environments, rather than”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    RSC responds strongly during passive viewing of environmental scenes in fMRI studies, especially when the scenes depict familiar places.

    “It responds strongly during passive viewing of environmental scenes in fMRI studies, especially when the scenes depict familiar places24.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [24] Epstein, R.A., Higgins, J.S., Jablonski, K. & Feiler, A.M. Visual scene processing in familiar and unfamiliar environments. J. Neurophysiol. 97, 3670–3683 (2007). source

      Not checked in the free run.

  • May combine two claims.

    Interpretationcitation

    A geometry-dependent but environment-independent representation of this type might be critical for translating viewpoint-dependent scene information into allocentric spatial codes-and, conversely, for re-creating viewpoint-dependent scenes from allocentric memory traces.

    “A geometry-dependent but environment-independent representation of this type might be critical for translating viewpoint-dependent scene information (”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [17] Byrne, P., Becker, S. & Burgess, N. Remembering the past and imagining the future: a neural model of spatial memory and imagery. Psychol. Rev. 114, 340–375 (2007). source

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    Locations were treated as equivalent across museums if they were in the equivalent position within the room, but not if they were in the equivalent position as defined by the global reference frame.

    “Moreover, locations were treated as equivalent across museums if they were in the equivalent position within the room, but not if they were in the equivalent position as defined by the global reference frame.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    Views facing the same local direction in different museums were treated as similar, as assessed by both priming and MVPA decoding, whereas views facing the same global direction were not.

    “Views facing the same local direction (as defined by local geometry) in different museums were treated as similar, as assessed by both priming and MVPA decoding, whereas views facing the same global direction were not.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationown data

    RSC represents imagined facing direction and imagined location during spatial memory recall; moreover, it does so using a reference frame that is anchored to local environmental features and generalizes across local environments with similar geometric structures.

    “Our results indicate that the retrosplenial complex (RSC), a region in human medial parietal lobe, represents imagined facing direction and imagined location during spatial memory recall; moreover, it does so using a reference frame that is anchored to local environmental features and generalizes across local environments with similar geometric structures.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    RSC response is increased when subjects retrieve spatial information, perform JRD tasks from memory, or move through an environment for which they have obtained survey knowledge.

    “Moreover, its response is increased when subjects retrieve spatial information25, 26, perform JRD tasks from memory27, or move through an environment for which they have obtained survey knowledge28.”
    Quote in the paperNot a substantial claim0 of 4 sources checked
    • cites [25] Epstein, R.A., Parker, W.E. & Feiler, A.M. Where am I now? Distinct roles for parahippocampal and retrosplenial cortices in place recognition. J. Neurosci. 27, 6141–6149 (2007). source

      Not checked in the free run.

    • cites [26] Rosenbaum, R.S., Ziegler, M., Winocur, G., Grady, C.L. & Moscovitch, M. “I have often walked down this street before”: fMRI studies on the hippocampus and other structures during mental navigation of an old environment. Hippocampus 14, 826–835 (2004). source

      Not checked in the free run.

    • cites [27] Zhang, H., Copara, M. & Ekstrom, A.D. Differential recruitment of brain networks following route and cartographic map learning of spatial environments. PLoS ONE 7, e44886 (2012). source

      Not checked in the free run.

    • cites [28] Wolbers, T. & Büchel, C. Dissociable retrosplenial and hippocampal contributions to successful formation of survey representations. J. Neurosci. 25, 3333–3340 (2005). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    RSC has been previously implicated in spatial navigation and spatial memory.

    “This region has been previously implicated in spatial navigation13, 22 and spatial memory14, 23.”
    Quote in the paperNot a substantial claim0 of 4 sources checked
    • cites [13] Epstein, R.A. Parahippocampal and retrosplenial contributions to human spatial navigation. Trends Cogn. Sci. 12, 388–396 (2008). source

      Not checked in the free run.

    • cites [22] Maguire, E.A. The retrosplenial contribution to human navigation: a review of lesion and neuroimaging findings. Scand. J. Psychol. 42, 225–238 (2001). source

      Not checked in the free run.

    • cites [14] Vann, S.D., Aggleton, J.P. & Maguire, E.A. What does the retrosplenial cortex do? Nat. Rev. Neurosci. 10, 792–802 (2009). source

      Not checked in the free run.

    • cites [23] Burgess, N. Spatial cognition and the brain. Ann. NY Acad. Sci. 1124, 77–97 (2008). source

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    The current results extend previous work by showing that RSC codes directions and locations while subjects perform a behavioral task that elicits direction-dependent and location-dependent effects.

    “The current results extend this previous work by showing that RSC codes directions and locations while subjects perform a behavioral task that elicits direction-dependent and location-dependent effects.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    When subjects perform a JRD task, they typically exhibit orientation-dependent performance.

    “When subjects perform a JRD task, they typically exhibit orientation-dependent (i.e. direction-dependent) performance.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [23] Burgess, N. Spatial cognition and the brain. Ann. NY Acad. Sci. 1124, 77–97 (2008). source

      Not checked in the free run.

    • cites [29] Burgess, N., Spiers, H.J. & Paleologou, E. Orientational manoeuvres in the dark: dissociating allocentric and egocentric influences on spatial memory. Cognition 94, 149–166 (2004). source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    In cases where subjects have learned multiple views of an environment, or have extensive experience over time, performance is typically best for a single imagined facing direction that acts as a "conceptual north," and it is better for facing directions that are aligned or orthogonal to this primary reference direction than for directions that are misaligned.

    “In cases where subjects have learned multiple views of an environment, or have extensive experience over time, performance is typically best for a single imagined facing direction that acts as a “conceptual north,” and it is better for facing directions that are aligned or orthogonal to this primary reference direction than for directions that are misaligned19, 20.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [19] Shelton, A.L. & McNamara, T.P. Systems of spatial reference in human memory. Cognit. Psychol. 43, 274–310 (2001). source

      Not checked in the free run.

    • cites [20] Mou, W. & McNamara, T.P. Intrinsic frames of reference in spatial memory. J. Exp. Psychol. Learn. Mem. Cogn. 28, 162–170 (2002). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    fMRI adaptation is observed in RSC when successively-presented scenes face the same direction.

    “fMRI adaptation is observed in RSC when successively-presented scenes face the same direction16”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [16] Baumann, O. & Mattingley, J.B. Medial parietal cortex encodes perceived heading direction in humans. J. Neurosci. 30, 12897–12901 (2010). source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation

    Previous work indicates that spatial geometry can be a dominant influence on the alignment of reference axes under many circumstances.

    “but previous work indicates that spatial geometry can be a dominant influence under many circumstances.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Geometry has long been known to guide reorientation, often to the detriment of other cues.

    “Indeed, geometry has long been known to guide reorientation, often to the detriment of other cues30, 31”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [30] Cheng, K. A purely geometric module in the rat's spatial representation. Cognition 23, 149–178 (1986). source

      Not checked in the free run.

    • cites [31] Gallistel, C.R. The Organization of Learning (The MIT Press, Cambridge, MA, 1990).

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation

    We previously reported that multivoxel patterns elicited in RSC when viewing familiar scenes contain information about location and direction that is independent of the specific view.

    “and we previously reported that multivoxel patterns elicited in RSC when viewing familiar scenes contain information about location and direction that is independent of the specific view4.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [4] Vass, L.K. & Epstein, R.A. Abstract representations of location and facing direction in the human brain. J. Neurosci. 33, 6133–6142 (2013). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    HD cells have been identified in the retrosplenial cortex of the rodent.

    “HD cells have been previously identified in the retrosplenial cortex of the rodent33, 34.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [33] Cho, J. & Sharp, P.E. Head direction, place, and movement correlates for cells in the rat retrosplenial cortex. Behav. Neurosci. 115, 3–25 (2001). source

      Not checked in the free run.

    • cites [34] Chen, L.L., Lin, L.-H., Green, E.J., Barnes, C.A. & McNaughton, B.L. Head-direction cells in the rat posterior cortex. Exp. Brain Res. 101, 8–23 (1994). source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    HD cells maintain a consistent sense of direction through path integration, referring back to the environment periodically to recalibrate when error has accumulated.

    “Rather, they maintain a consistent sense of direction through path integration37, only referring back to the environment periodically to recalibrate when error has accumulated10.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [37] McNaughton, B.L., Battaglia, F.P., Jensen, O., Moser, E.I. & Moser, M.-B. Path integration and the neural basis of the 'cognitive map'. Nat. Rev. Neurosci. 7, 663–678 (2006). source

      Not checked in the free run.

    • cites [10] Valerio, S. & Taube, J.S. Path integration: how the head direction signal maintains and corrects spatial orientation. Nat. Neurosci. 15, 1445–1453 (2012). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    There is striking overlap among brain regions involved in retrieval of autobiographical memories, imagination of novel events, and navigation.

    “Much recent interest has focused on the striking overlap among brain regions involved in the retrieval of autobiographical memories, the imagination of novel events, and navigation14, 47, 48.”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites [14] Vann, S.D., Aggleton, J.P. & Maguire, E.A. What does the retrosplenial cortex do? Nat. Rev. Neurosci. 10, 792–802 (2009). source

      Not checked in the free run.

    • cites [47] Ranganath, C. & Ritchey, M. Two cortical systems for memory-guided behaviour. Nat. Rev. Neurosci. 13, 713–726 (2012). source

      Not checked in the free run.

    • cites [48] Hassabis, D., Kumaran, D. & Maguire, E.A. Using imagination to understand the neural basis of episodic memory. J. Neurosci. 27, 14365–14374 (2007). source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation

    Hippocampal place cells fired in analogous positions in geometrically identical environments learned in a single training session, even with proprioceptive and vestibular cues present.

    “similar results were observed in a recent rodent neurophysiology study which found that hippocampal place cells fired in analogous positions in geometrically identical environments that were learned in a single training session, even though these proprioceptive and vestibular cues were present45.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [45] Spiers, H.J., Hayman, R.M.A., Jovalekic, A., Marozzi, E. & Jeffery, K.J. Place field repetition and purely local remapping in a multicompartment environment. Cereb. Cortex 10.1093/cercor/bht198 (13 August 2013). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    These regions, including RSC, form a network that mediates both prospective thought and episodic memory.

    “These regions, including RSC, are believed to form a network that mediates both prospective thought and episodic memory49.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [49] Schacter, D.L. et al. The future of memory: remembering, imagining, and the brain. Neuron 76, 677–694 (2012). source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    RSC is strongly activated when imagining changes in one's viewpoint or when retrieving spatial relationships following first-person navigation.

    “This conclusion is consistent with previous work indicating that RSC is strongly activated when imagining changes in one’s viewpoint50 or when retrieving spatial relationships following first-person navigation27.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [50] Lambrey, S., Doeller, C., Berthoz, A. & Burgess, N. Imagining being somewhere else: neural basis of changing perspective in space. Cereb. Cortex 22, 166–174 (2012). source

      Not checked in the free run.

    • cites [27] Zhang, H., Copara, M. & Ekstrom, A.D. Differential recruitment of brain networks following route and cartographic map learning of spatial environments. PLoS ONE 7, e44886 (2012). source

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    Subjects knew where the object was within the spatial frame but did not remember the museum to which the frame should be applied.

    “This suggests that they knew where the object was within the spatial frame, but did not remember the museum to which the frame should be applied.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    RSC anchors our sense of direction to the world, allowing us to stay oriented while navigating and to reconstruct from memory the experience of being in a particular place.

    “Like a compass rose, RSC anchors our sense of direction to the world, thus allowing us to stay oriented while we navigate and to reconstruct from memory the experience of being in a particular place.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductnone given

    Sample size was determined based on a pilot behavioral experiment.

    “Sample size was determined based on a pilot behavioral experiment.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    Six additional participants from Experiment 2 were excluded prior to scanning for scoring less than 70% on the behavioral practice trials after the first day of training.

    “Six additional participants from Experiment 2 were excluded prior to scanning for scoring less than 70% on the behavioral practice trials after the first day of training”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Reference frames can be observed across many different scales, from tabletops to rooms to campuses.

    “Previous behavioral work with non-nested environments has indicated that reference frames can be observed across many different scales, from tabletops, to rooms19, to campuses43.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [19] Shelton, A.L. & McNamara, T.P. Systems of spatial reference in human memory. Cognit. Psychol. 43, 274–310 (2001). source

      Not checked in the free run.

    • cites [43] Marchette, S.A., Yerramsetti, A., Burns, T.J. & Shelton, A.L. Spatial memory in the real world: long-term representations of everyday environments. Mem. Cognit. 39, 1401–1408 (2011). source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    In Experiment 2, assignment of objects to alcoves was randomized across participants to ensure decoding of direction and location could not be attributed to object identity.

    “In Experiment 2, we randomized the assignment of objects to alcoves in the environment across participants, to ensure that reliable decoding of direction and location could not be attributed to decoding of object identity.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    Three additional subjects scanned in Experiment 2 were excluded before analysis for technical difficulties in fMRI acquisition, early termination, and sleeping.

    “Three additional subjects were scanned in Experiment 2 but were excluded before analysis: the first for technical difficulties in fMRI acquisition, the second because she requested to terminate the scanning session early, and the third for sleeping.”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    Transition probabilities were adopted to ensure sufficient trials to detect priming of direction within a given museum.

    “These transition probabilities were adopted to ensure we would have sufficient trials to detect priming of direction within a given museum.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    The placement of museums dissociated direction within a museum (local North) from direction in the exterior park (global North).

    “This had the effect of dissociating direction within a museum as defined by the axis of entry and internal geometry (e.g. facing the back wall, or local North) from direction in the exterior park (e.g. facing the mountains, or global North).”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductreasoning

    The method for defining RSC ensures RSC can be defined in both hemispheres for every subject.

    “This method ensures that RSC can be defined in both hemispheres for every subject.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).

    Study design and conductreasoning

    The longer interval between response and subsequent trial in Experiment 2 might have reduced behavioral priming effects.

    “Note that this meant that there was a substantially longer interval between a participants’ response and the subsequent trial, which might have reduced behavioral priming effects.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductown data

    For four subjects, the 16th view was not represented within the continuous carryover sequence due to technical errors, and analysis was adjusted to exclude this view.

    “For four subjects, the 16th view was not represented within the continuous carryover sequence due to technical errors. For these subjects, the analysis was performed as normal but adjusted to exclude this view.”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim.

    Study design and conductown data

    The MVPA procedure allowed testing whether information carried by patterns in the first half of the experiment generalized to patterns in the second half.

    “This procedure allowed us to test whether the information carried by patterns of responses in the first half of the experiment (i.e. the first carryover sequence) generalized to patterns in the second half.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim.

    Study design and conductcitation

    Monte Carlo simulations permuting the sign of whole-brain maps were performed to find the true type I error rate.

    “To find the true type I error rate, we performed Monte Carlo simulations that permute the sign of the whole-brain maps from individual subjects58.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [58] Nichols, T.E. & Holmes, A.P. Nonparametric permutation tests for functional neuroimaging: a primer with examples. Hum. Brain Mapp. 15, 1–25 (2002). source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Study design and conductnone given

    Only voxels surviving correction for multiple comparisons across the entire brain at p < 0.05 were reported.

    “We reported only the voxels that survive correction for multiple comparisons across the entire brain, significant at p < 0.05.”
    Quote in the paperSubstantial claim
67 of 67

  • The paper relies on this without support.

    A neural locus for a separate representation of locally-referenced facing direction has not yet been identified.

    “Although there is considerable evidence that both rodents and humans use allothetic information to orient themselves, a neural locus for this separate representation of locally-referenced facing direction has yet to be identified.”

  • The authors state this as an assumption.

    Data distributions were assumed to be normal, but this was not formally tested.

    “Data distributions were assumed to be normal, but this was not formally tested.”
2 of 2

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

  • In participants who learned object locations in geometrically similar but visually distinct virtual buildings, the representation of facing direction was generalized across environments with similar geometry, as indicated by fMRI activity pattern similarity in the RSC across multiple trials.

    “In participants who learned the location of objects in geometrically similar but visually distinct (virtual) buildings 152 , the representation of facing direction (for example, south-west) was generalized across environments with similar geometry, as indicated by fMRI activity pattern similarity in the RSC across multiple trials.”

    From From cognitive maps to spatial schemas (2023) · cites this paper as [152]

  • The RSC integrates egocentric and allocentric spatial frames of reference and generalizes local codes across geometrically similar environments.

    “The RSC has also been shown to integrate egocentric and allocentric spatial frames of reference 146,152 and to generalize local codes (that is, codes for specific regions, such as the southwest corner of a specific room) across geometrically similar environments (in this example, to the south-west corner of various rooms) 19,39,152 .”

    From From cognitive maps to spatial schemas (2023) · cites this paper as [152]

Statistics that don’t add up

Each reported p-value recomputed from its test statistic and degrees of freedom.

  • Inconsistent p-value: t(44) = 4.599, p = .00008

    Minor

    The paper reports p = .00008 here; recomputing from the test statistic and its degrees of freedom gives p < .001 (two-tailed). The check cannot tell whether the statistic or the p-value is the misprint.

    Reported p = .00008 · recomputed p < .001

1 of 1

Statistics recomputed

Every reported test result, recomputed from its test statistic and degrees of freedom. A result shows as not adding up only once an independent check has confirmed it.

  • Doesn’t add upt(44) = 4.599reported p = .00008 · recomputed p < .001

    “t(44) = 4.599, p = .00008”

  • Not confirmedF(1, 23) = 8.279reported p = .008 · recomputed p = .009

    “F(1,23) =8.279, p = .008”

  • ConsistentF(1, 21) = 11.631reported p = .003 · recomputed p = .003

    “F(1,21) = 11.631, p = .003”

  • ConsistentF(1, 21) = 11.802reported p = .002 · recomputed p = .002

    “F(1,21) = 11.802, p = .002”

  • ConsistentF(1, 21) = 0.734reported p = .401 · recomputed p = .401

    “F(1,21) = 0.734, p = .401”

  • ConsistentF(1, 21) = 5.695reported p = .026 · recomputed p = .027

    “F(1,21) = 5.695, p = .026”

  • ConsistentF(1, 21) = 4.227reported p = .052 · recomputed p = .052

    “F(1,21) = 4.227, p = .052”

  • ConsistentF(1, 21) = 0.012reported p = .915 · recomputed p = .914

    “F(1,21) = 0.012, p = .915”

  • Consistentt(21) = 1.283reported p = .214 · recomputed p = .213

    “t(21) = 1.283, p = .214”

  • Consistentt(21) = 0.976reported p = .34 · recomputed p = .340

    “t(21) = 0.976, p = .340”

  • Consistentt(21) = 4.242reported p = .00036 · recomputed p < .001

    “t(21) = 4.242, p = .00036”

  • ConsistentF(1, 23) = 1.501reported p = .233 · recomputed p = .233

    “F(1,23) = 1.501, p = .233”

  • ConsistentF(1, 23) = 1.19reported p = .287 · recomputed p = .287

    “F(1,23) = 1.190, p = .287”

  • ConsistentF(1, 23) = 20.009reported p = .00017 · recomputed p < .001

    “F(1,23) = 20.009, p = .00017”

  • ConsistentF(1, 23) = 2.45reported p = .131 · recomputed p = .131

    “F(1,23) = 2.450, p = .131”

  • ConsistentF(1, 23) = 0.005reported p = .942 · recomputed p = .944

    “F(1,23) = 0.005, p = .942”

  • ConsistentF(1, 23) = 0.85reported p = .366 · recomputed p = .366

    “F(1,23) = 0.850, p = .366”

  • ConsistentF(1, 23) = 1.043reported p = .318 · recomputed p = .318

    “F(1,23) = 1.043, p = .318”

  • ConsistentF(1, 23) = 7.162reported p = .013 · recomputed p = .013

    “F(1,23) = 7.162, p = .013”

  • ConsistentF(1, 23) = 2.719reported p = .113 · recomputed p = .113

    “F(1,23) = 2.719, p = .113”

  • ConsistentF(1, 23) = 0.284reported p = .6 · recomputed p = .599

    “F(1,23) = 0.284, p = .600”

  • ConsistentF(1, 23) = 4.581reported p = .043 · recomputed p = .043

    “F(1,23) = 4.581, p = .043”

  • ConsistentF(1, 23) = 1.834reported p = .189 · recomputed p = .189

    “F(1,23) = 1.834, p = .189”

  • ConsistentF(1, 23) = 0.768reported p = .39 · recomputed p = .390

    “F(1,23) = 0.768, p = .390”

  • Consistentt(23) = 0.724reported p = .476 · recomputed p = .476

    “t(23) = 0.724, p = .476”

  • Consistentt(23) = 0.965reported p = .344 · recomputed p = .345

    “t(23) = 0.965, p = .344”

  • Consistentt(23) = 3.495reported p = .002 · recomputed p = .002

    “t(23) = 3.495, p = .002”

  • ConsistentF(2, 46) = 6.675reported p = .003 · recomputed p = .003

    “F(2,46) = 6.675, p = .003”

  • ConsistentF(1, 23) = 3.2reported p = .087 · recomputed p = .087

    “F(1,23) = 3.200, p = .087”

  • ConsistentF(2, 46) = 0.316reported p = .73 · recomputed p = .731

    “F(2,46) = 0.316, p = .730”

  • ConsistentF(1, 23) = 8.615reported p = .007 · recomputed p = .007

    “F(1,23) = 8.615, p = .007”

  • Consistentt(23) = 2.652reported p = .014 · recomputed p = .014

    “t(23) = 2.652, p = .014”

32 of 32

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

  • Carryover sequence has more trials than a once-per-pair design permits

    Minor

    With 16 views, having each view precede every other view, including itself, exactly once requires 256 ordered transitions. Testing each view 17 times gives 272 trials per sequence and at least 271 transitions, so some ordered pairs must occur more than once.

1 of 1

Cites

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Cited by

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Related

Authors
Marchette, S. A., Vass, L. K., Ryan, J., & Epstein, R. A.
Affiliations
University of Pennsylvania (US)
Corresponding author
Marchette, Steven A
ORCID
Venue
Nature Neuroscience
Publisher
Springer Science and Business Media LLC
Published
2014-10-05
Volume
17
Issue
11
Pages
1598-1606
ISSN
  • 1097-6256 (ISSN-L)
  • 1546-1726
DOI
10.1038/nn.3834
PMC
PMC4309016
OpenAlex
W2082940604
Semantic Scholar
67a2da8324383ab894046aed172bd36edcecdfc4
Cited by
287
References
61
Citations by year
2026: 17 · 2025: 19 · 2024: 19 · 2023: 23 · 2022: 16 · 2021: 30 · 2020: 18 · 2019: 25 · 2018: 35 · 2017: 28 · 2016: 33 · 2015: 22 · 2014: 2
FWCI
34.69
Type
journal article
Language
en
Update notices
erratum (2015-05-26)
Open access
Licence
Locations
  • Nature Neuroscience (publishedVersion)
  • PubMed (publishedVersion)
  • PubMed Central (acceptedVersion)
Funders
National Eye Institute (R01EY022350, EY022350)
Topics
  • Physical Sciences › Engineering › Automotive Engineering › Spatial Cognition and Navigation
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
  • Life Sciences › Neuroscience › Neurology › Vestibular and auditory disorders
Keywords
retrosplenial complex, spatial reference frames, head direction, spatial navigation, landmark-based navigation, spatial memory, virtual worlds, multi-voxel pattern analysis, behavioral priming
MeSH terms
Adolescent, Adult, Brain Mapping, Female, Humans, Image Processing, Computer-Assisted, Learning, Magnetic Resonance Imaging, Male, Orientation, Parietal Lobe, Mental Recall, Space Perception, Spatial Behavior, Young Adult

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-09.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-09.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

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